【文献メモ】海洋における交雑(Gardner, 1997):未完 | ウッカリカサゴのブログ

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Jonathan P. A. Gardner (1997)
Hybridization in the Sea
海洋における交雑
Pages 1-78, in ADVANCES IN MARINE BIOLOGY VOL. 31
https://www.sciencedirect.com/bookseries/advances-in-marine-biology/vol/31/suppl/C

https://www.researchgate.net/publication/251449465_Hybridization_in_the_Sea

 

1. はじめに
   1.1. 背景と定義
   1.2. レビューの範囲

2. 交雑の認識
   2.1. 理論的考察
   2.2. 実際的な考慮事項
3. 分類学の重要性
   3.1. 系統学、分類学、および交雑
   3.2. 種の概念と交雑
4. 海洋での交雑を阻害または促進する要因
   4.1. 交雑を妨げる要因
   4.2. 交雑を促進する要因
5. 海洋での交雑はどのくらいの頻度で行われるか?
6. 交雑と繁殖のタイミング
7. 実験的な交雑
8. 交雑速度の時間的変動と安定性
9. 撹乱と交雑
   9.1. 導入
   9.2. 人為的妨害
   9.3. 自然災害
10. 親族の豊かさと交雑
11. 交雑個体の適合度
12. 生殖生物学と交雑頻度
13. 交雑の構造
   13.1. 交雑ゾーン幅
   13.2. 交雑ゾーンの形状
14. 遺伝子移入
15. 交雑バイアス: 存在するか?
16. 属間交雑と種間交雑
17. 海洋での交雑の生物地理学
  17.1. 海洋での交雑の地理的分布
  17.2. 生態学的差別化
18. 化石の記録
19. 漁業と保全の問題としての交雑
20. 海洋における交雑と種分化
21. 結論と今後の展望
表1
謝辞
参考文献



アブスクラクト


 この章では海における交雑(雑種化)について説明する。このレビューの主な目的は次のとおりである。
(1) 海での交雑はこれまで考えられていたほど稀ではないことを指摘すること。
(2) 海洋生物の特定のグループ間の交雑バイアスに注意を喚起すること。
(3) 交雑場所における生物地理学的パターンと環境パターンを説明すること。
(4) 海洋での交雑と他の環境での交雑を比較すること。
(5) 水産科学と保全の文脈で海洋での交雑について議論すること。
 この章では、環境に関係なく、交雑の 5つのレベルについて説明する。海における自然交雑を詳述した事例の概要が、交雑事象の地理的位置および生物地理的領域に関する情報とともに提供される。タンパク質電気泳動は、雑種や混合祖先の個体を識別できる特に有用なツールであることが証明されている。この技術により、2つの交雑分類群が 1つまたは複数のアロザイム遺伝子座における電気泳動の違いについて固定されている場合、個体の交雑状態の客観的な評価が可能になる。

 この章では、交雑ゾーンの定義において分類法が重要な役割を果たすことについて説明する。海洋交雑を妨げる要因と促進する要因についても議論する。実験室での交雑実験は、機会があれば 2つの分類群が交雑すること、または 2つの分類群からの配偶子が人工的に交雑できることを実証することにより、特定の個体のおそらく雑種の性質を確認するために広く利用されてきた。このような情報は、海洋での自然交雑の研究に貴重な貢献を提供する。

1. はじめに
   1.1. 背景と定義


 交雑と交雑ゾーンは、受精前または接合後のメカニズムによる生殖隔離に寄与する種分化とプロセスを研究する優れた機会を提供するため、進化生物学者にとって長い間興味深いものであった。
 このようなプロセスには、同類交雑、雑種不適合、配偶子不和合などが含まれ、これらすべてが種分化に寄与すると考えられている。

 おそらく驚くべきことかもしれないが、「交雑ゾーン」という用語は、定義が難しいことが判明している。定義には、すべての場合に適切であるとは限らないゾーンの起源や維持に関する基礎的なメカニズムが含まれたり暗示されたりすることが多いためである(たとえば、ゾーンの幅に関する推測など) またはゾーン内の個体の適合度)。
 最も実行可能な定義は、Harrison(1990, p. 72) によって提案されたものである。交雑ゾーンとは、

「少なくともいくつかの混合祖先の子孫をもたらす、遺伝的に異なる個体グループ間の相互作用である。遺伝的に異なる 2つのグループの純粋な集団は、相互作用ゾーンの外側で見つかる。」 

これは、このレビューで採用されている交雑ゾーンの定義である。
 同様に、雑種形成の孤立した例は、遺伝的に異なる 2つの親個体の交雑から生じる 1つまたは複数の個体の生産として定義できる。
 これからわかるように、海中における孤立した雑種形成の頻度は、交雑ゾーンや群れ(後者は、遺伝的に異なる親個体が遺伝子移入を伴って高度に交雑した結果、混血した個体の集まりと定義される)の存在頻度よりもかなり高い。

 過去10年間、交雑ゾーンは大きな注目を集めてきた(たとえば、Endler, 1977; Moore, 1977; Barton and Hewitt, 1985, 1989; Harrison, 1990; Arnold, 1992; Arnold, 1993) によるレビュー)。
 さまざまなタイプの交雑ゾーン(クライン、テンション ゾーン、モザイク、網状など) とそのようなゾーンの確立と維持に関与する要因については十分に文書化されており、読者はこれらの優れたレビューを参照することができる。
 この論文の目的は、これらのアイデアや概念を再検討することではなく、これまでほとんど注目されてこなかった海洋における自然交雑に関する新しい情報を提供することである。

 上で引用したレビューの中で、海中での自然交雑について言及しているのは 1 人の著者だけであり、十分に文書化された事例はほとんど存在しないと述べられているのは、興味深いと同時に驚くべきことである(Harrison、1993、p. 111)。
 陸上の自然交雑についてはさらに多くのことが知られており、陸上環境からより多くのより適切に文書化された例が存在することは事実であるが、海での自然交雑は珍しい現象ではないことも事実である。
 明らかに、海洋での交雑を調べた研究からのデータである。海は推定体積 137 x 10 6 km3 であり、地球上で最大の生態系であり(Gage と Tyler、1991)、地球の 71% 以上をカバーしている(Briggs、1991)、そして多くのユニークな門(May, 1988; Winston, 1992) に貢献するこの論文は、自然交雑とその原因と結果についての私たちの理解を実質的に助けるに違いない。

 交雑は、種分化における最も重要なステップの 1つとみなされることが多い。なぜなら、交雑する分類群が融合しない場合、それらの分類群は、交雑相手の選択の差異および/または配偶子不適合性などのメカニズムを進化させて雑種形成を妨げ、それによって種分化を促進すると考えられるからである(e.g. Mayr, 1942) 。
 海洋環境の文脈では、種分化については比較的ほとんど知られていない(Knox、1963; Palumbi、1992、1994)。
 海洋環境は継続的であり(Battaglia、1957; Sara、1985)、他の環境に比べて一時的ではない(Hubbs and Kuronuma、1941)。

 さらに、陸上システムで見られる比較的顕著なエコトーンの種類は、海洋環境ではそれほど頻繁ではない。
 多くの海洋種は広範囲に分散する可能性があるため、空間的に分離された個体群間の遺伝的差異は、多くの場合わずかである(例、Skibinski et al., 1983)。
 しかし、生態学的不均一性(例えば、潮間帯や汽水域)は、(微)地理的隔離と組み合わされて、海洋環境における適応分化と最終的には進化的分岐を促進するはずであることが指摘されている(Battaglia、1957)。
 Harrison(1990) の定義によれば、遺伝的分化は交雑と交雑ゾーン形成の前提条件であるため、これは交雑を促進する可能性がある。

 海における自然交雑に関する異質で、時には不明瞭なデータを調べると、交雑のプロセス自体をより深く理解するために、また海における種分化についてのより深い洞察を得るために説明しなければならない多くのパターンが存在することが示されている。
 このレビューの主な目的は次のとおりである。
(1) 海中での交雑はこれまで考えられていたほど稀ではないことを指摘すること。
 (2) 海洋生物の特定のグループ間の交雑バイアスに注意を喚起すること。
 (3) 交雑の場所における生物地理学的パターンと環境パターンを説明すること。
 (4) 海洋での交雑と他の環境での交雑を比較すること。
 (5) 水産科学と保全の文脈で海洋での交雑について議論すること。


   1.2. レビューの範囲

 このレビューでは、海洋での交雑の例をできるだけ多く含めることを可能にするために、比較的広範なアプローチが採用されている。
 例は、完全な海洋から河口までの範囲の環境条件から抽出される。
 河口域での交雑の包含は、次の理由により正当化される。
(1) 河口域は、多くの真の海洋種にとって、脱皮、生殖、摂食、幼生の育苗場などとして生態学的に重要である。
(2) これらの地域は海洋の種分化を促進する上で重要な役割を果たしている。
 遡河性(サケ科など) と降潮性(ウナギなど) の両方の種が関与する交雑の例は除外される。そのような動物は淡水で繁殖するか、生涯のほとんどを淡水で過ごすため、真の海洋生物と同じ進化圧力を受けないからである。生物。
 レビューの範囲は、単一の(通常は不妊の)個体の生産を伴う自然交雑の孤立した例から、高レベルの遺伝子移入を伴う海洋雑種群の事例、染色体の倍加を伴う種分化の事例まで多岐にわたる。
 私が知っている限り多くの例を取り上げたが、このレビューはすべてを網羅したものではない。
主題のせいで(雑種形成の可能性のある孤立した例は「出版可能性」の尺度ではそれほど高くない)、この情報の多くは不明瞭であり、読者はここに含まれていない例に精通しているかもしれない。

 私が意図しているのは、海洋交雑の概要、そのパターン、偏り、影響を把握するために、できるだけ多くの事例から引き出すことである。限られた事例では、文献に矛盾する報告があるため、自然交雑が本当に起こっているかどうかを判断するのは難しい。
 例えば、ザルガイ Cerastodenna edule と C. glaucum は北西ヨーロッパで同所的に発生し(Brock, 1979; Gosling, 1980)、これら 2つの優れた種の間の形態学的中間にある個体は自然雑種であることが示唆されている(Kingston, 1973)。
 他の場所では、交雑がこの形態学的仲介の説明ではないことを示唆する証拠がある(Brock, 1978; Gosling, 1980)。
 このような場合、交雑の証拠がいくつかあるにもかかわらず、証拠の大部分がこれら 2つの種の間で自然交雑が起こらないことを示しているため、この例はレビューに含まれない。

 交雑の発生の可能性に関する見解の相違のより最近の例には、サバ科のヨコシマサワラ  Scomberomorus commerson と タイワンサワラ S. guttatus が含まれる。
 Srinivasa Rao と Lakshmi(1993) によると、これら 2 種はインド南東海岸沖で交雑し、以前は S. lineolatus として認識されていた雑種を生み出した。Collette(1994) はこの解釈に強く反対しているが、双方が提示したデータは生化学的(アロザイム変異など) や分子的(DNA 配列データ) ではなく骨学的なものであるため、どちらの議論の妥当性を判断するか困難である。
 この例は、どちらの意見にも重きを置くことなく、完全を期すためにこのレビューに含まれている。最後に、海洋交雑記録の多くには、海洋交雑ゾーンの存在の証拠のない孤立した交雑例が含まれていることは明らかである。
 これらの記録は、これらのデータに貴重な補足を提供するため、より文書化された交雑ゾーンおよび交雑スウォームの事例とともにレビューに含まれている。

 

以下、ここでは省略

 

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表1
Table 1 Summary table of recent hybridization in the sea._

Biogeographic zones: 
  CT, cold temperate 
  P, polar 
  T, tropical 
  WT, warm temperate

Characters examined: 
  A, allozymes 
  B, behaviour 
  C, colour 
  K, karyotype 
  M, morphology 
  MH, molecular hybridization 
  Mt, mitochondrial DNA 
  N, nuclear DNA

Disturbance: 
  H, human 
  S, salinity 
  U, upwelling 
  Y, yes (some other cause)

References: (■魚類)
  1 Alma$a (1972) 
■2 Ayling (1980) 
  3 Bates (1992) 
  4 Beaumont et al. (1989) 
  5 Bert (1986) 
  6 Bert and Harrison (1988) 
  7 Bert et al. (1993) 
  8 Boss (1964)
  9 Brown (1995) 
  10 Burrows and Lodge (1951) 
  11 Chia (1966) 
■12 Cocks (1887) 
  13 Coustau et al. (1991a) 
  14 Coustau et al. (1991b) 
  15 Coyer and Zaugg-Haglund (1982) 
  16 Coyer et al. (1992) 
  17 Crane (1975) 
■18 Daget (1963) 
■19 Dahlberg (1969a) 
■20 Dahlberg (1969b) 
■21 Dahlberg (1970) 
■22 Devaraj (1986) 
  23 Dillon (1992) 
  24 Dillon and Manzi (1989) 
■25 Duggins et al. (1995) 
■26 Dymond (1939) 
  27 Edwards and Skibinski (1987) 
■28 Feddern (1968) 
■29 Fischer (1980) 
  30 Fujino et al. (1980) 
■31 Fujio (1977) 
  32 Gardner and Skibinski (1988) 
  33 Gardner and Skibinski (1990a) 
  34 Gardner and Skibinski (1990b) 
  35 Gardner and Skibinski (1991a) 
  36 Gardner and Skibinski (1991b) 
  37 Gardner and Thompson (unpubl.) 
  38 Gartner-Kepkay et al. (1980) 
  39 Gartner-Kepkay et al. (1983) 
■40 Gosline (1948) 
  41 Gosling and Wilkins (1981) 
  42 Grant et al. (1977) 
  43 Gray et al. (1991) 
  44 Hagstrom and Lonning (1961) 
  45 Hagstrom and Lonning (1964) 
■46 Hart (1973) 
  47 Hedgecock et al. (1993) 
■48 Herald (1941) 
  49 Hesselman et al. (1988) 
■50 Hettier (1968) 
■51 Holt (1893) 
■52 Hubbs and Kuronuma (1941) 
  53 Johnson (1976) 
  54 Jones and Naylor (1971) 
  55 Karinen and Hoopes (1971) 
  56 Knight and Parke (1950) 
  57 Koehn et al. (1984) 
  58 Kwast et al. (1990) 
  59 Liu et al. (1991) 
■60 McClure and McEachran (1992) 
  61 McDonald and Koehn (1988) 
  62 McDonald et al. (1991) 
  63 Menge (1986) 
  64 Menzel (1985) 
  65 Menzel and Menzel (1965) 
  66 Miller (1994) 
  67 Miller and Babcock (unpubl.) 
  68 Miller and Benzie (unpubl.) 
  69 Neushul (1962) 
■70 Norman (1934) 
  71 Owen et al. (1971) 
  72 Porter and Chestnut (I960) 
■73 Pyle and Randall (1994) 
■74 Randall and Fridman (1981) 
■75 Randall et al. (1977) 
■76 Rivas (I960) 
■77 Russell (1988) 
■78 Sale (1991) 
  79 Sarver and Foltz (1993) 
  80 Sarver et al. (1992) 
  81 Sasaki et al. (1980) 
  82 Schilder (1962)
  83 Schopf and Murphy (1973) 
■84 Schultz and Smith (1936) 
  85 Skibinski (1983) 
  86 Skibinski (1985) 
  87 Solignac (1976) 
  88 Solignac (1981) 
■89 Spilliaert et al, (1991) 
■90 Srinivasa Rao and Ganapati (1977) 
■91 Srinivasa Rao and Lakshmi (1993) 
  92 Strathmann (1981) 
  93 Swan (1953) 
  94 Swan (1962) 
  95 Talmadge (1977) 
  96 Tokida et al. (1958) 
■97 Turner (1967) 
■98 Turner (1969) 
  99 Vainola and Hvilsom (1991) 
 100 Vainola and Varvio (1989) 
 101 Varvio et al. (1988) 
 102 Vasseur (1952) 
 103 Verrill (1909) 
 104 Victor (1986) 
 105 Wilbur (1989) 
 106 Willis and Skibinski (1992)

Superscripts: 
  A, the authors suggest hybrid unfitness (ref. 6) 
  B, the author suggests no hybrid unfitness (ref. 103) 
  C, hybrids have lower wet weight (but intermediate size) and greater morphological variability than parents (ref. 58) 
  D, laboratory studies indicate no abnormalities among hybrids (ref. 11) 
  E, high incidence of neoplasia in hybrids (ref. 49) 
  F, hybrids have increased growth rate and better shelf life than parents (ref. 64) 
  G, hybrids exhibit intermediate rates of parasitism (ref. 19) 
  H, hybrids exhibit intermediate fat reserves (ref. 20) 
  I, hybrids observed to spawn (ref. 21) 
  J, sex ratio of hybrids dominated by males (ref. 21) 
  K, ref. 2 
  L, ref. 75 
  M, ref. 2 
  N, ref. 76 
  O, ref. 82 
  P, ref. 48 
  Q, ref. 22 
  R, hybrids, on average, greater length than at least one parent (ref. 12) 
  *, these species economically important as shellfish predators 
  **, these hybrids mentioned but not described 
  +, now known as H. bermudensis (ref. 71)